Kugadzirwa kwemupunga kuri kudzikira nekuda kwekushanduka kwemamiriro ekunze uye kusiyanisa muColombia.Plant growth regulatorsyakashandiswa senzira yekudzikisa kupisa kunetseka muzvirimwa zvakasiyana. Naizvozvo, chinangwa chechidzidzo ichi chaive chekuongorora maitiro emuviri (stomatal conductance, stomatal conductance, yakazara chlorophyll content, Fv/Fm ratio yemaviri ekutengesa mupunga genotypes akaiswa mukusangana kwekupisa (kupisa kwemasikati nehusiku), tembiricha yecanopy uye yehuwandu hwemvura) uye biochemical variables (malondialdehyde (MDA) uye prolinic acid content). Yekutanga neyechipiri kuyedza yakaitwa pachishandiswa zvirimwa zviviri zvemupunga genotypes Federrose 67 ("F67") uye Federrose 2000 ("F2000"), zvichiteerana. Zvose zviedzo zvakaongororwa pamwe chete senhevedzano yezviedzo. Mishonga yakadzikwa yaitevera: kudzora zvachose (AC) (zvinomera zvemupunga zvinokura patembiricha yakanyatsonaka (tembiricha yemasikati/usiku 30/25°C)), kudzora kushushikana kwekupisa (SC) [zvinomera mupunga zvakasangana nekushushikana kwekupisa chete (40/25°C). 30 ° C)], uye zvirimwa zvemupunga zvakasimbiswa uye zvakapfapfaidzwa nemagadzirirwo ekukura kwemiti (stress + AUX, stress + BR, stress + CK kana stress + GA) kaviri (5 mazuva asati uye 5 mazuva mushure mekupisa kwekupisa). Kupfapfaidza neSA kwakawedzera huwandu hwechlorophyll yemhando mbiri (huremu hutsva hwemiti yemupunga "F67" uye "F2000" yaive 3.25 uye 3.65 mg/g, zvichiteerana) zvichienzaniswa nezvirimwa zveSC (huremu hutsva hwe "F67" zvirimwa hwaive 2.36 uye 2.56 mg) mupunga "arFlix" uye 0CK application ye 0,0CK zvakare ye 0. kazhinji yakavandudza stomatal conductance yemupunga "F2000" zvirimwa (499.25 vs. 150.60 mmol m-2 s) zvichienzaniswa nekudzora kushushikana kwekupisa. kushushikana kwekupisa, kudziya kwekorona yechirimwa kunoderera ne2-3 ° C, uye iyo MDA yemukati mumiti inoderera. Chiyero chekutsungirira chinoratidza kuti kushandiswa kwefoliar kweCK (97.69%) uye BR (60.73%) kunogona kubatsira kuderedza dambudziko rekupisa kwakabatanidzwa. kushushikana zvakanyanya muF2000 mupunga zvirimwa. Mukupedzisa, kupfapfaidza foliar kweBR kana CK kunogona kutorwa seagronomic zano rekubatsira kudzikisira mhedzisiro yekusangana kwekushushikana kwekupisa pahunhu hwehupenyu hwezvirimwa zvemupunga.
Mupunga (Oryza sativa) ndewemhuri yePoaceae uye ndeimwe yezviyo zvinorimwa zvakanyanya pasi rose pamwe nechibage negorosi (Bajaj naMohanty, 2005). Nzvimbo inorimwa mupunga mahekita 617,934, uye kugadzirwa kwenyika muna 2020 kwaive matani 2,937,840 neavhareji goho re5.02 matani/ha (Federarroz (Federación Nacional de Arroceros), 2021).
Kudziya kwepasirese kuri kukanganisa zvirimwa zvemupunga, zvichitungamira kumhando dzakasiyana dzekushushikana kweabiotic sekupisa kwakanyanya uye nguva yekusanaya kwemvura. Kuchinja kwemamiriro ekunze kuri kuita kuti kupisa kwepasi rose kukwire; Tembiricha inofungidzirwa kuti ichakwira ne1.0–3.7°C muzana ramakore rechi21, izvo zvinogona kuwedzera kuwanda uye kusimba kwekushushikana kwekupisa. Kuwedzera kwekushisa kwezvakatipoteredza kwakanganisa mupunga, zvichiita kuti goho ridzikire ne6-7%. Nekune rimwe divi, kushanduka kwemamiriro ekunze kunotungamirawo kumamiriro ekunze asina kunaka ezvirimwa, senge nguva yekusanaya kwemvura zvakanyanya kana kupisa kwakanyanya munzvimbo dzinopisa uye dzinopisa . Pamusoro pezvo, zviitiko zvekusiyana zvakaita seEl Niño zvinogona kukonzera kushushikana kwekupisa uye kuwedzera kukuvadzwa kwechirimwa mune mamwe matunhu anopisa. MuColombia, tembiricha munzvimbo dzinogadzira mupunga dzinofungidzirwa kuti dzichawedzera ne2–2.5°C panosvika 2050, zvichideredza kugadzirwa kwemupunga uye kukanganisa kuyerera kwechigadzirwa kumisika nemaketani ekutengesa.
Zvizhinji zvirimwa zvemupunga zvinorimwa munzvimbo dzine tembiricha dziri pedyo nepamusoro pekukura kwembesa (Shah et al., 2011). Zvinonzi optimal avhareji masikati neusiku tembiricha nokudamupunga kukura uye kukuraanowanzo 28 ° C uye 22 ° C, zvichiteerana (Kilasi et al., 2018; Calderón-Páez et al., 2021). Tembiricha dziri pamusoro pezvikumbaridzo izvi zvinogona kukonzera nguva dzepakati kusvika kune dzakanyanya kushushikana kwekupisa panguva dzakaoma nhanho dzekukura kwemupunga (kuchera, kupikisa, maruva, uye kuzadza zviyo), zvichikanganisa goho rezviyo. Uku kuderedzwa kwegoho kunonyanya kukonzerwa kwenguva yakareba yekupisa kupisa, iyo inokanganisa plant physiology . Nekuda kwekudyidzana kwezvinhu zvakasiyana-siyana, senge kushushikana kwenguva uye yakanyanya tembiricha yasvika, kupisa kushushikana kunogona kukonzera kuwanda kwekukuvara kusingadzoreki pakudyara metabolism nekusimudzira.
Kushushikana kwekupisa kunokanganisa akasiyana physiological uye biochemical maitiro muzvirimwa. Leaf photosynthesis ndeimwe yemaitiro anonyanya kunetseka nekupisa muzvirimwa zvemupunga, sezvo mwero wephotosynthesis unodzikira ne50% apo tembiricha yezuva nezuva inodarika 35°C. Mapinduriro emuviri wezvirimwa zvemupunga anosiyana zvichienderana nerudzi rwekushushikana kwekupisa. Semuyenzaniso, photosynthetic rates uye stomatal conductance zvinodzivirirwa kana zvirimwa zvakasangana nekupisa kwemasikati kukuru (33–40°C) kana tembiricha yepamusoro masikati nehusiku (35–40°C masikati, 28–30°C). C zvinoreva husiku) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Kupisa kwehusiku kwepamusoro (30 ° C) kunokonzera kudzivirirwa zvine mwero kwephotosynthesis asi kuwedzera kufema kwehusiku (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Pasinei nenguva yekunetseka, kupisa kwekushisa kunobatawo mashizha e chlorophyll content, chiyero che chlorophyll variable fluorescence kusvika kune yakanyanya chlorophyll fluorescence (Fv / Fm), uye Rubisco activation mumiti yemupunga (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso et al., 2014).
Kuchinja kwebiochemical chimwe chikamu chekugadzirisa kwezvirimwa kune kupisa kunetseka (Wahid et al., 2007). Proline yemukati yakashandiswa sechiratidzo chebiochemical chekushushikana kwechirimwa (Ahmed naHassan 2011). Proline inobata basa rinokosha mumiti yemasabolism sezvo inoshanda se carbon kana nitrogen sosi uye se membrane stabilizer pasi pemamiriro ekupisa kwepamusoro (Sánchez-Reinoso et al., 2014). Kupisa kwepamusoro kunobatawo kugadzikana kwe membrane kuburikidza ne lipid peroxidation, zvichiita kuti pave nekuumbwa kwemalondialdehyde (MDA) (Wahid et al., 2007). Nokudaro, zvinyorwa zveMDA zvakashandiswawo kunzwisisa kutendeseka kwemaitiro emagetsi emagetsi pasi pekupisa kwekupisa (Cao et al., 2009; Chavez-Arias et al., 2018). Pakupedzisira, kusanganiswa kwekupisa kupisa [37/30 ° C (masikati / usiku)] kwakawedzera chikamu che electrolyte leakage uye malondialdehyde zviri mumupunga (Liu et al., 2013).
Kushandiswa kwezvinomera kukura regulators (GRs) kwakaongororwa kuti kuderedze migumisiro yakaipa yekupisa kupisa, sezvo zvinhu izvi zviri kushingaira kubatanidzwa mumhinduro dzemiti kana nzira dzekudzivirira muviri pakurwisana kwakadaro (Peleg naBlumwald, 2011; Yin et al., 2011; Ahmed et al., 2015). Exogenous kushandiswa kwemajini zviwanikwa kwakave nemhedzisiro yakanaka pakupisa kusagadzikana kushivirira muzvirimwa zvakasiyana. Zvidzidzo zvakaratidza kuti phytohormones yakadai segibberellins (GA), cytokinins (CK), auxins (AUX) kana brassinosteroids (BR) inotungamirira kuwedzera kwezvakasiyana-siyana zvehupenyu uye biochemical variables (Peleg naBlumwald, 2011; Yin et al. Ren, 2011; Mitler, Z201; Mitler et al. 2014). MuColombia, kushandiswa kwekunze kwezviwanikwa zvemajini uye maitiro ayo pazvirimwa zvemupunga hazvina kunyatsonzwisiswa nekudzidzwa. Zvisinei, chidzidzo chekare chakaratidza kuti foliar spraying ye BR inogona kuvandudza kushivirira kwemupunga nekuvandudza gasi kuchinjana maitiro, chlorophyll kana proline zvinyorwa zvemashizha emupunga (Quintero-Calderón et al., 2021).
Cytokinins inopindirana mhinduro dzezvirimwa kune abiotic kusagadzikana, kusanganisira kupisa kunetseka (Ha et al., 2012). Mukuwedzera, zvakataurwa kuti kushandiswa kwekunze kweCK kunogona kuderedza kukuvara kwekupisa. Semuenzaniso, kushandiswa kwekunze kwezeatin kwakawedzera chiyero chephotosynthetic, chlorophyll a uye b zvemukati, uye erekitironi yekufambisa zvakanaka mune zvinokambaira bentgrass (Agrotis estolonifera) panguva yekupisa kunetseka (Xu naHuang, 2009; Jespersen naHuang, 2015). Exogenous application ye zeatin inogonawo kuvandudza antioxidant basa, kuwedzera kuumbwa kwemapuroteni akasiyana-siyana, kuderedza reactive oxygen marudzi (ROS) kukuvadza uye malondialdehyde (MDA) kugadzirwa mumiti yemiti (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar et al., 2020).
Kushandiswa kwegibberellic acid kwakaratidzawo mhinduro yakanaka kune kupisa kunetseka. Zvidzidzo zvakaratidza kuti GA biosynthesis inopindirana nzira dzakasiyana-siyana dzemagetsi uye inowedzera kushivirira pasi pemamiriro ekushisa kwepamusoro (Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi et al. (2020) akawana kuti kupfapfaidza foliar kweiyo exogenous GA (25 kana 50 mg * L) inogona kuwedzera photosynthetic chiyero uye antioxidant chiitiko mukupisa-kusimbisa zvirimwa zveorenji zvichienzaniswa nezvinomera zvinodzora. Izvo zvakaonekwa zvakare kuti exogenous kushandiswa kweHA kunowedzera hunyoro hunyoro, chlorophyll uye carotenoid zviri mukati uye inoderedza lipid peroxidation mumiti yemichindwe (Phoenix dactylifera) pasi pekupisa kunetseka (Khan et al., 2020). Auxin inotorawo basa rakakosha mukugadzirisa mhinduro dzekukura kune yakakwirira tembiricha mamiriro (Sun et al., 2012; Wang et al., 2016). Uyu mutongi wekukura anoshanda semucherechedzo webiochemical mumaitiro akasiyana-siyana akadai seproline synthesis kana kuderedzwa pasi pekunetseka kweabiotic (Ali et al. 2007). Pamusoro pezvo, AUX inowedzerawo antioxidant basa, izvo zvinotungamira mukudzikira kweMDA muzvirimwa nekuda kwekuderera kwe lipid peroxidation (Bielach et al., 2017). Sergeev nevamwe. (2018) akacherechedza kuti mumiti yepea (Pisum sativum) pasi pekupisa kunetseka, zvinyorwa zveproline - dimethylaminoethoxycarbonylmethyl) naphthylchloromethyl ether (TA-14) inowedzera. Mukuyedza kumwe chete, vakacherekedza mazinga akaderera eMDA mumiti yakarapwa zvichienzaniswa nezvirimwa zvisina kurapwa neAUX.
Brassinosteroids imwe kirasi yekukura regulators inoshandiswa kudzikisira mhedzisiro yekupisa kusagadzikana. Ogweno et al. (2008) yakashuma kuti exogenous BR spray yakawedzera mambure photosynthetic rate, stomatal conductance uye chiyero chikuru cheRubisco carboxylation yematomatisi (Solanum lycopersicum) zvinomera pasi pekupisa kwemazuva masere. Foliar spraying ye epibrassinosteroids inogona kuwedzera mambure photosynthetic mwero wegaka (Cucumis sativus) zvinomera pasi pekupisa kunetseka (Yu et al., 2004). Uye zvakare, exogenous kushandiswa kweBR kunonoka kushatisa chlorophyll uye kunowedzera kushandiswa kwemvura kunyatsoshanda uye yakawanda quantum goho rePSII photochemistry muzvirimwa pasi pekupisa kunetseka (Holá et al., 2010; Toussagunpanit et al., 2015).
Nekuda kwekushanduka kwemamiriro ekunze uye kushanduka, zvirimwa zvemupunga zvinotarisana nenguva dzekupisa kwemazuva ese (Lesk et al., 2016; Garcés, 2020; Federarroz (Federación Nacional de Arroceros), 2021). Mumiti inonzi phenotyping, kushandiswa kwe phytonutrients kana biostimulants kwakadzidzwa senzira yekuderedza kupisa kwemhepo munzvimbo dzinorimwa mupunga (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Mukuwedzera, kushandiswa kwezvinyorwa zvebiochemical uye physiological variables (kupisa kwemashizha, stomatal conductance, chlorophyll fluorescence parameters, chlorophyll uye hukama hwemvura, malondialdehyde uye proline synthesis) chinhu chakavimbika chekuongorora zvirimwa zvemupunga pasi pekupisa kwekupisa munyika uye kune dzimwe nyika (Sánchez -Reynoso et al., Al. 2017; Zvisinei, tsvakurudzo pamusoro pekushandiswa kwefoliar phytohormonal sprays mumupunga wenharaunda inoramba isingawanzoitiki, saka, kuongorora kwemaitiro ehupenyu uye maitiro ekugadzirwa kwezvirimwa zvakakosha zvikuru kune chirevo chemaitiro agronomic ekugadzirisa izvi. chlorophyll fluorescence paramita uye inoenderana mvura yemukati) uye biochemical mhedzisiro yefoliar kushandiswa kwechina chirimwa kukura regulators (AUX, CK, GA uye BR). (Photosynthetic pigments, malondialdehyde uye proline contents) Kusiyana mune maviri ekutengesa mupunga genotypes akaiswa pasi pekupisa kwakabatanidzwa (kupisa kwemasikati / usiku).
Muchidzidzo ichi, zviedzo zviviri zvakazvimiririra zvakaitwa. The genotypes Federrose 67 (F67: genotype yakagadzirwa mukupisa kwakanyanya mukati memakore gumi apfuura) uye Federrose 2000 (F2000: genotype yakagadzirwa mumakore gumi ekupedzisira ezana remakore rechi20 ichiratidza kurwisana nehutachiona hwemashizha machena) yakashandiswa kekutanga. mbeu. uye kuyedza kwechipiri, zvichiteerana. Ose maviri genotypes anorimwa zvakanyanya nevarimi vekuColombia. Mbeu dzakadyarwa mu10-L matireyi (kureba 39.6 cm, hupamhi 28.8 cm, kureba 16.8 cm) iine ivhu rejecha rine 2% organic matter. Mhodzi shanu dzisati dzamera dzakadyarwa mutireyi yega yega. Mapallet akaiswa mugreenhouse yeFaculty of Agricultural Sciences yeNational University of Colombia, Bogotá campus (43°50′56″ N, 74°04′051″ W), panzvimbo yakakwirira ye2556 m pamusoro pegungwa (asl). m.) uye zvakaitwa kubva muna Gumiguru kusvika Zvita 2019. Imwe kuyedza (Federroz 67) uye yechipiri kuyedza (Federroz 2000) mumwaka mumwe chete we2020.
Mamiriro ezvakatipoteredza mugreenhouse mumwaka wega wega wekudyara ndeaya: masikati neusiku tembiricha 30/25°C, humidity 60~80%, natural photoperiod maawa gumi nemaviri (photosynthetically active radiation 1500 µmol (photons) m-2 s-). 1 masikati). Zvirimwa zvakaiswa fetiraiza zvinoenderana nezviri mukati mechinhu chimwe nechimwe mazuva makumi maviri mushure mekubuda kwembeu (DAE), maererano naSánchez-Reinoso et al. (2019): 670 mg nitrogen pachirimwa, 110 mg phosphorus pachirimwa, 350 mg potassium pachirimwa, 68 mg calcium pachirimwa, 20 mg magnesium pachirimwa, 20 mg sarufa pachirimwa, 17 mg silicon pachirimwa. Miti yacho ine 10 mg boron pachirimwa, 17 mg copper pachirimwa, uye 44 mg zinc pachirimwa. Zvidyarwa zvemupunga zvakachengetwa kusvika ku47 DAE mukuedza kwega kwega pavakasvika phenological stage V5 panguva iyi. Zvidzidzo zvekare zvakaratidza kuti iyi nhanho ye phenological inguva yakakodzera yekuita zvidzidzo zvekupisa kupisa mupunga (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Mukuedza kwega kwega, mashandisirwo maviri akasiyana emugadziri wekukura kwemashizha akaitwa. Iyo yekutanga seti yefoliar phytohormone sprays yakashandiswa mazuva mashanu isati yasvika kupisa kwekushushikana kwekurapa (42 DAE) kugadzirira zvirimwa zvekunetseka kwezvakatipoteredza. Yechipiri yekupfapfaidza yefoliar yakazopihwa mazuva mashanu mushure mekunge zvirimwa zvave pachena kune stress mamiriro (52 DAE). Zvine phytohormones zvakashandiswa uye zvinhu zvechimwe nechimwe chinoshandiswa chakapfapfaidzwa muchidzidzo ichi chakanyorwa muSupplementary Table 1. Mamiriro ekugadzirisa mashizha ekukura kwemashizha akashandiswa akaita seaya: (i) Auxin (1-naphthylacetic acid: NAA) pamusangano we5 × 10-5 M (ii) 5 × 5 × gibberelli acid (5 × gibberelli) gibberelli (5 × gibberelli) gibberelli; GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. Aya maitiro akasarudzwa nokuti anokonzera mhinduro dzakanaka uye anowedzera kumira kwemiti kune kupisa kunetseka (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Rice zvirimwa pasina chero chirimwa kukura regulator sprays aibatwa nemvura distilled chete. Miti yese yemupunga yakapfapfaidzwa nechipupa chemaoko. Isa 20 ml H2O pachirimwa kuti unyorovese nzvimbo dzepamusoro nedzezasi dzemashizha. Masprayers ese efoliar akashandisa adjuvant yekurima (Agrotin, Bayer CropScience, Colombia) pa 0.1% (v/v). Nhambwe iri pakati pepoto nesprayer i30cm.
Kurapa kwekushushikana kwekupisa kwaiitwa mazuva mashanu mushure mekutanga foliar spray (47 DAE) mukuedza kwega kwega. Zvidyarwa zvemupunga zvakatamiswa kubva mugreenhouse kuenda ku294 L yekukura mukamuri (MLR-351H, Sanyo, IL, USA) kumisikidza kushushikana kwekupisa kana kuchengetedza zvakafanana mamiriro ezvakatipoteredza (47 DAE). Kurapa kwekushushikana kwekupisa kwakasanganiswa kwakaitwa nekuisa kamuri kune tembiricha yezuva/usiku inotevera: tembiricha yepamusoro yemasikati [40°C kwemaawa mashanu (kubva pa11:00 kusvika 16:00)] uye nhambo yeusiku [30°C kwemaawa mashanu]. Mazuva masere akateedzana (kubva 19:00 kusvika 24:00). Kushushikana kwekushisa uye nguva yekuratidzira yakasarudzwa zvichienderana nezvidzidzo zvekare (Sánchez-Reynoso et al. 2014; Alvarado-Sanabría et al. 2017). Kune rumwe rutivi, boka remiti inotamirwa kukamuri yekukura yakachengetwa mu greenhouse pane imwe tembiricha (30 ° C masikati / 25 ° C usiku) kwemazuva masere akatevedzana.
Pakupera kwekuedza, mapoka ekurapa anotevera akawanikwa: (i) kukura kwekushisa kwemamiriro ezvinhu + kushandiswa kwemvura yakasvibiswa [Absolute control (AC)], (ii) kupisa kwemamiriro ekushisa + kushandiswa kwemvura yakasvibiswa [Heat stress control (SC)], (iii) mamiriro ekupisa kwemamiriro ezvinhu + auxin application (AUX), (iv) kupisa kushushikana kwemamiriro ezvinhu + gibberellin application (GAVI +TOCK), (vtoCK kupisa kwemhepo), (vtoCK kupisa) + brassinosteroid (BR) Appendix. Aya mapoka ekurapa akashandiswa kune maviri genotypes (F67 uye F2000). Ese marapirwo akaitwa mune yakasarudzika dhizaini ine zvinyorwa zvishanu, imwe neimwe iine chirimwa chimwe. Chirimwa chimwe nechimwe chakashandiswa kuverenga zvakasiyana-siyana zvakatemwa pakupera kwekuedza. Kuedza kwakatora 55 DAE.
Stomatal conductance (gs) yakayerwa pachishandiswa porosometer inotakurika (SC-1, METER Group Inc., USA) inotangira pa0 kusvika 1000 mmol m-2 s-1, ine sample chamber aperture ye6.35 mm. Zviyero zvinotorwa nekubatanidza stomameter probe kune shizha rakura uye pfuti huru yechirimwa yakawedzera zvizere. Pakurapa kwega kwega, kuverenga kwegs kwakatorwa pamashizha matatu echirimwa chimwe nechimwe pakati pe11:00 na16:00 uye avhareji.
RWC yakatemwa maererano nenzira yakatsanangurwa naGhoulam et al. (2002). Iro pepa rakawedzerwa zvizere rinoshandiswa kuona g rakashandiswawo kuyera RWC. Huremu hutsva (FW) hwakatarwa pakarepo mushure mekukohwa pachishandiswa chiyero chedigital. Mashizha acho aizoiswa mumudziyo wepurasitiki wakazadzwa nemvura ndokusiiwa murima panzvimbo yetembiricha (22°C) kwemaawa 48. Wobva wayera pachiyero chedhijitari uye nyora huremu hwakawedzerwa (TW). Mashizha akazvimba akaomeswa pa75 ° C kwemaawa makumi mana nemasere uye huremu hwavo hwakaoma (DW) hwakanyorwa.
Hukama hwe chlorophyll hwakatemerwa uchishandisa mita yechlorophyll (paLeafmeter, FT Green LLC, USA) uye inoratidzwa muzvikamu zveLeaf (Dey et al., 2016). PSII yakanyanya kuwanda yekuverenga kuverenga (Fv/Fm ratio) yakarekodhwa vachishandisa inoenderera mberi yekunakidzwa chlorophyll fluorimeter (Handy PEA, Hansatech Instruments, UK). Mashizha aive akasviba-achichinjika achishandisa mashizha masimbi e20 min pamberi peFv/Fm zviyero (Restrepo-Diaz uye Garces-Varon, 2013). Mushure mokunge mashizha akanga akasviba acclimated, baseline (F0) uye yakanyanya fluorescence (Fm) yakayerwa. Kubva pane idzi data, variable fluorescence (Fv = Fm - F0), chiyero che fluorescence yakasiyana kusvika kune yakanyanya fluorescence (Fv / Fm), iyo yakawanda yehuwandu yegoho rePSII photochemistry (Fv / F0) uye reshiyo Fm / F0 yakaverengwa (Baker, 2008; Lee et al., 2017). Relative chlorophyll uye chlorophyll fluorescence kuverenga kwakatorwa pamashizha mamwe anoshandiswa kuyera gs.
Inenge 800 mg yemashizha matsva uremu yakaunganidzwa se biochemical variables. Sampuli dzemashizha dzakazoiswa homogenized mu nitrogen yemvura uye dzakachengetwa kuti dziwedzere kuongororwa. Iyo spectrometric nzira inoshandiswa kufungidzira tishu chlorophyll a, b uye carotenoid zvirimo zvinobva pane nzira uye equations inotsanangurwa naWellburn (1994). Masampuli emashizha emashizha (30 mg) akaunganidzwa uye akaiswa homogenized mu 3 ml ye80% acetone. Iwo masampuli akabva aiswa centrifuged (modhi 420101, Becton Dickinson Primary Care Diagnostics, USA) pa5000 rpm kwemaminetsi gumi kubvisa zvidimbu. Iyo supernatant yakaderedzwa kusvika kuvhoriyamu yekupedzisira ye6 ml nekuwedzera 80% acetone (Sims naGamon, 2002). Zviri muchlorophyll zvakatemwa pa663 (chlorophyll a) uye 646 (chlorophyll b) nm, uye carotenoids pa470 nm vachishandisa spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Iyo thiobarbituric acid (TBA) nzira inotsanangurwa naHodges et al. (1999) yakashandiswa kuongorora membrane lipid peroxidation (MDA). Inenge 0.3 g yemashizha emashizha akaiswawo homogenized mu nitrogen yemvura. Iwo masampula aive centrifuged pa5000 rpm uye kunyura kwakayerwa pane spectrophotometer pa440, 532 uye 600 nm. Pakupedzisira, kusungirirwa kweMDA kwakaverengerwa uchishandisa kupera kwekofi (157 M mL-1).
Proline yezvinyorwa zvese zvekurapa zvakatemwa uchishandisa nzira yakatsanangurwa naBates et al. (1973). Wedzera 10 ml ye 3% aqueous solution ye sulfosalicylic acid kumuenzaniso wakachengetwa uye sefa kuburikidza neWhatman filter paper (Nhamba. 2). Zvadaro 2 ml yefiltrate iyi yakabatwa ne 2 ml ye ninhydric acid uye 2 ml yeglacial acetic acid. Iko kusanganiswa kwakaiswa mumvura yekugeza pa 90 ° C kweawa imwe. Misa kuita nekuisa paaizi. Zunza chubhu nesimba uchishandisa vortex shaker uye nyungudutsa mhinduro mu 4 ml ye toluene. Kuverenga kweAbsorbance kwakatemwa pa520 nm vachishandisa iyo yakafanana spectrophotometer yakashandiswa kuverengera photosynthetic pigments (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Nzira yakatsanangurwa naGerhards et al. (2016) kuverenga tembiricha yecanopy uye CSI. Thermal mifananidzo yakatorwa neFLIR 2 kamera (FLIR Systems Inc., Boston, MA, USA) nekururama kwe ± 2 ° C pakupera kwekunetseka. Isa nzvimbo yakachena kuseri kwechirimwa kuti utore mifananidzo. Zvakare, mafekitari maviri aionekwa semienzaniso yereferensi. Miti yacho yakaiswa panzvimbo yakachena; imwe yaive yakashongedzwa nemubatsiri wezvekurima (Agrotin, Bayer CropScience, Bogotá, Colombia) kutevedzera kuvhurwa kwese stomata [wet mode (Twet)], uye imwe yaive shizha pasina chero application [Dry mode (Tdry)] (Castro -Duque et al., 2020). Nharaunda pakati pekamera nepoto panguva yekutorwa kwemafirimu yaive 1 m.
Indekisi yekushivirira yakaenzana yakaverengerwa zvisina kunanga uchishandisa stomatal conductance (gs) yemiti yakarapwa kana ichienzaniswa nekudzora zvirimwa (zvinomera zvisina kunetsekana kurapwa uye nekukura regulators vakaiswa) kuona kushivirira kweakarapwa genotype akaongororwa muchidzidzo ichi. RTI yakawanikwa uchishandisa equation yakagadziriswa kubva kuChávez-Arias et al. (2020).
Muchiyedzo chega chega, zvese zvakasiyana-siyana zvemuviri zvataurwa pamusoro zvakatemwa uye kurekodhwa pa55 DAE vachishandisa mashizha akawedzerwa akaunganidzwa kubva padenga repamusoro. Mukuwedzera, zviyero zvakaitwa mukamuri yekukura kuti zvirege kuchinja mamiriro ezvakatipoteredza umo zvinomera zvinokura.
Data kubva kuongororo yekutanga neyechipiri yakaongororwa pamwe chete seyakatevedzana yekuedza. Boka rega rega rekuyedza raive nezvirimwa zvishanu, uye chirimwa chega chega chaive chikamu chekuyedza. Kuongororwa kwekusiyana (ANOVA) kwakaitwa (P ≤ 0.05). Apo misiyano yakakura yakaonekwa, Tukey's post hoc comparative test yakashandiswa paP ≤ 0.05. Shandisa iyo arcsine basa kushandura muzana maitiro. Data yakaongororwa pachishandiswa Statistix v 9.0 software (Analytical Software, Tallahassee, FL, USA) uye yakarongwa kushandisa SigmaPlot (version 10.0; Systat Software, San Jose, CA, USA). Chikamu chikuru chekuongorora chakaitwa pachishandiswa InfoStat 2016 software (Analysis Software, National University of Cordoba, Argentina) kuti ione zvakanakisisa zvinomera zvinomera pasi pekudzidza.
Tafura 1 inopfupisa iyo ANOVA inoratidza zviedzo, marapirwo akasiyana, uye kudyidzana kwavo nemashizha photosynthetic pigments (chlorophyll a, b, yakazara, uye carotenoids), malondialdehyde (MDA) uye proline content, uye stomatal conductance. Mhedzisiro ye gs, hukama hwemvura. (RWC), chlorophyll content, chlorophyll alpha fluorescence parameters, korona tembiricha (PCT) (°C), chirimwa stress index (CSI) uye hama yekushivirira index yezvirimwa zvemupunga pa55 DAE.
Tafura 1. Pfupiso ye ANOVA data pamusoro pemupunga physiological uye biochemical variables pakati pekuedza (genotypes) uye kupisa kunetseka kurapwa.
Kusiyana (P≤0.01) mumashizha e photosynthetic pigment interactions, zvinyorwa zve chlorophyll (Atleaf readings), uye alpha-chlorophyll fluorescence parameters pakati pekuedza uye kurapwa kunoratidzwa muTebhu 2. Kupisa kwemasikati uye kwehusiku kwakawedzera huwandu hweklorophyll uye zvinyorwa zvecarotenoid. Mbeu dzemupunga dzisina kana foliar spray ye phytohormones (2.36 mg g-1 ye "F67" uye 2.56 mg g-1 ye "F2000") zvichienzaniswa nezvinomera zvinorimwa pasi pemamiriro ekushisa akanyanya (2.67 mg g -1)) yakaratidza kuderera kweklorophyll. Mune zvese zviedzo, "F67" yaive 2.80 mg g-1 uye "F2000" yaive 2.80 mg g-1. Pamusoro pezvo, madzinde emupunga akabatwa nemusanganiswa weAUX neGA sprays pasi pekupisa kwekupisa kwakaratidzawo kuderera kwehuwandu hwechlorophyll mune ese genotypes (AUX = 1.96 mg g-1 uye GA = 1.45 mg g-1 ye "F67" ; AUX = 1.96 mg g-1 uye GA "F6 g-7" ye "F6" GA = 1. 2.24 mg) g-1 uye GA = 1.43 mg g-1 (ye "F2000" ) pasi pemamiriro ekupisa. Pasi pemamiriro ekupisa kwekupisa, kurapwa kwefoliar neBR kwakakonzera kuwedzera kudiki kwekuchinja uku mune ese genotypes. Pakupedzisira, CK foliar spray yakaratidza yakakwira photosynthetic pigment kukosha pakati pese marapiro (AUX, GA, BR, SC uye AC marapirwo) mu genotypes F67 (3.24 mg g-1) uye F2000 (3.65 mg g-1). Izvo zvine hushamwari zve chlorophyll (Atleaf unit) zvakare yakaderedzwa nekusanganiswa kupisa kusagadzikana. Hunhu hwepamusoro hwakanyorwa zvakare muzvirimwa zvakapfapfaidzwa neCC mune ese genotypes (41.66 ye "F67" uye 49.30 ye "F2000"). Fv neFv/Fm ratidzo dzakaratidza kusiyana kukuru pakati pemishonga nemacultivars (Table 2). Pakazere, pakati peizvi zvakasiyana, cultivar F67 yakanga isinganyanyi kunetseka nekupisa kupfuura cultivar F2000. Mareshiyo eFv neFv/Fm akatambura zvakanyanya mukuyedza kwechipiri. Yakasimbiswa 'F2000′ mbesa dzisina kupfapfaidzwa chero phytohormones dzaive neiyo yakaderera Fv kukosha (2120.15) uye Fv/Fm ratios (0.59), asi foliar kumwaya neCK kwakabatsira kudzoreredza hunhu uhu (Fv: 2591, 89, Fv/Fm 3 chiyero: 0. , kugamuchira kuverenga kwakafanana nekwakanyorwa pamiti ye "F2000" yakakura pasi pemamiriro ekushisa akakodzera (Fv: 2955.35, Fv / Fm ratio: 0.73: 0.72). Pakanga pasina misiyano yakakosha mukutanga fluorescence (F0), yakanyanya fluorescence (Fm), yakanyanya photochemical quantum goho rePSII (Fv / F0) uye Fm / F0 chiyero. Pakupedzisira, BR yakaratidza maitiro akafanana sezvakaonekwa neCK (Fv 2545.06, Fv/Fm chiyero 0.73).
Tafura 2. Mhedzisiro yekubatana kwekupisa kunetseka (40 ° / 30 ° C masikati / usiku) pamashizha photosynthetic pigments [total chlorophyll (Chl Total), chlorophyll a (Chl a), chlorophyll b (Chl b) uye carotenoids Cx + c] effect ], jamaa chlorophyll unit), resiniophyll parameter unit (Atiniophyll). fluorescence (F0), yakanyanya fluorescence (Fm), variable fluorescence (Fv), yakanyanya PSII kunyatsoshanda (Fv/Fm), photochemical maximum quantum goho rePSII (Fv/F0) uye Fm/F0 muzvirimwa zvemupunga maviri genotypes [Federrose 67 (F67) uye Federrose 00 mushure me50 mazuva 20 (DAE)).
Relative water content (RWC) yemiti yemupunga yakasiyana-siyana yakaratidza misiyano (P ≤ 0.05) mukubatana pakati pekuedza uye kurapwa kwefoliar (Fig. 1A). Kana yakarapwa neSA, iyo yakaderera kukosha yakanyorwa kune ese ari maviri genotypes (74.01% yeF67 uye 76.6% yeF2000). Pasi pemamiriro ekupisa kwekupisa, iyo RWC yezvirimwa zvemupunga zveese genotypes akabatwa neakasiyana phytohormones akawedzera zvakanyanya. Pakazere, kushandiswa kwefoliar kweCK, GA, AUX, kana BR kwakawedzera RWC kune hunhu hwakafanana nehwezvinomera zvinokura pasi pemamiriro akanyanya panguva yekuyedza. Kudzora kwakazara uye foliar yakapfapfaidza zvidyarwa zvakarekodha kukosha kweanosvika 83% kune ese ari maviri genotypes. Kune rumwe rutivi, gs yakaratidzawo kusiyana kwakasiyana (P ≤ 0.01) mukuedza-kurapa kusangana (Fig. 1B). Iyo absolute control (AC) chirimwa zvakare chakarekodha epamusoro kukosha kune yega genotype (440.65 mmol m-2s-1 yeF67 uye 511.02 mmol m-2s-1 yeF2000). Zvidyarwa zvemupunga zvakasangana nekushushikana kwekupisa kwega kwakaratidza hutsika hwakaderera hwema genotypes (150.60 mmol m-2s-1 yeF67 uye 171.32 mmol m-2s-1 yeF2000). Foliar kurapwa pamwe zvose zvinomera kukura regulators yakawedzerawo g. PaF2000 mupunga zvirimwa zvakapfapfaidzwa neCC, mhedzisiro yekupfapfaidza foliar ne phytohormones yaive pachena. Iri boka rezvirimwa rakaratidza kusasiyana kana zvichienzaniswa neabsolute control plant (AC 511.02 uye CC 499.25 mmol m-2s-1).
Mufananidzo 1. Mhedzisiro yekubatana kwekupisa kunetseka (40 ° / 30 ° C masikati / usiku) pahuwandu hwemvura (RWC) (A), stomatal conductance (gs) (B), malondialdehyde (MDA) kugadzirwa (C), uye proline content. (D) muzvirimwa zvemupunga maviri genotypes (F67 uye F2000) pamazuva makumi mashanu neshanu mushure mekubuda (DAE). Mishonga yakaongororwa yejenotype yega yega yaisanganisira: kudzora zvachose (AC), kupisa kudzvinyirira (SC), kupisa kushushikana + auxin (AUX), kupisa kunetseka + gibberellin (GA), kupisa kunetseka + cell mitogen (CK), uye kupisa kunetseka + brassinosteroid. (BR). Koramu yega yega inomiririra iyo ± yakajairwa kukanganisa kwema data mashanu mapoinzi (n = 5). Makolamu anoteverwa nemabhii akasiyana anoratidza mutsauko wakakosha maererano nebvunzo yaTukey (P ≤ 0.05). Mavara ane chiratidzo chakaenzana anoratidza kuti chirevo hachina kukosha (≤ 0.05).
MDA (P ≤ 0.01) uye proline (P ≤ 0.01) zvinyorwa zvakaratidzawo kusiyana kukuru mukubatana pakati pekuedza uye phytohormone mishonga (Fig. 1C, D). Kuwedzera lipid peroxidation akaonekwa pamwe SC kurapwa mune zvose genotypes (Figure 1C), zvisinei zvirimwa kurapwa shizha kukura regulator pfapfaidzo yakaratidza yakaderera lipid peroxidation mune zvose genotypes; Kazhinji, kushandiswa kwe phytohormones (CA, AUC, BR kana GA) kunotungamirira mukuderera kwe lipid peroxidation (MDA content). Hapana misiyano yakawanikwa pakati pe AC zvirimwa zviviri genotypes uye zvirimwa pasi pekunetseka kwekupisa uye yakapfapfaidzwa ne phytohormones (yakacherechedzwa FW kukosha mu "F67" zvidyarwa zvakabva pa4.38-6.77 µmol g-1, uye muFW "F2000" zvirimwa "zvakacherechedzwa tsika kubva ku92.18 kusvika 18 g imwe 2. ruoko, proline synthesis mu "F67" zvidyarwa zvaive zvakaderera pane "F2000" zvirimwa pasi pekunetsekana kwakabatanidzwa, izvo zvakaita kuti kuwedzera kwekugadzirwa kweproline mukupisa-kusimbiswa kwezvirimwa zvemupunga, mune zvose zviedzo, zvakaonekwa kuti kutungamirirwa kwehomoni idzi kwakawedzera zvakanyanya amino acid yemiti yeF2000 (AUX uye 43l4 g-30) kuremekedza. (Mufananidzo 1G).
Mhedzisiro yefoliar plant growth regulator spray uye yakasanganiswa kupisa kushushikana pachirimwa checanopy tembiricha uye hama yekushivirira index (RTI) inoratidzwa muFigure 2A uye B. Kune ese genotypes, tembiricha yecanopy yezvirimwa zve AC yaive pedyo ne27 ° C, uye iyo yeSC zvirimwa yaive yakapoteredza 28 ° C. WITH. Zvakaonekwawo kuti kurapwa kwefoliar neCK neBR kwakakonzera kudzikira kwe2–3°C mukupisa kwecanopy zvichienzaniswa nezvinomera zveSC (Mufananidzo 2A). RTI yakaratidza maitiro akafanana kune mamwe maitiro ehupenyu, kuratidza kusiyana kukuru (P ≤ 0.01) mukubatana pakati pekuedza uye kurapwa (Mufananidzo 2B). Zvidyarwa zveSC zvakaratidza kushivirira kwechirimwa mune ese genotypes (34.18% uye 33.52% ye "F67" uye "F2000" zvirimwa zvemupunga, zvichiteerana). Kudyisa foliar ye phytohormones inovandudza RTI muzvirimwa zvakafumurwa kune yakanyanya tembiricha kusagadzikana. Izvi zvainyanya kutaurwa mu "F2000" zvidyarwa zvakapfapfaidzwa neCC, umo iyo RTI yaive 97.69. Kune rumwe rutivi, misiyano yakakura yakaonekwa chete mugoho rekusimbisa indekisi (CSI) yemiti yemupunga pasi pefoliar factor spray stress mamiriro (P ≤ 0.01) (Fig. 2B). Chete zvidyarwa zvemupunga zvakaiswa pasi pekunetseka kwekupisa kwakaomarara kwakaratidza iyo yakanyanya kushushikana index kukosha (0.816). Kana zvidyarwa zvemupunga zvakapfapfaidzwa neakasiyana phytohormones, index yekushushikana yaive yakaderera (tsika kubva 0.6 kusvika 0.67). Pakupedzisira, chirimwa chemupunga chakakura pasi pemamiriro akanaka chaive nemutengo we0.138.
Mufananidzo 2. Migumisiro yekubatana kwekupisa kwekupisa (40 ° / 30 ° C masikati / usiku) pane tembiricha yekanopi (A), indekisi yekushivirira (RTI) (B), uye index stress index (CSI) (C) yemarudzi maviri emiti. Commercial mupunga genotypes (F67 uye F2000) vakaiswa pasi siyana kupisa kurapwa. Mishonga yakaongororwa yejenotype yega yega yaisanganisira: kudzora zvachose (AC), kupisa kudzvinyirira (SC), kupisa kushushikana + auxin (AUX), kupisa kunetseka + gibberellin (GA), kupisa kunetseka + cell mitogen (CK), uye kupisa kunetseka + brassinosteroid. (BR). Kushushikana kwekupisa kwakasanganiswa kunosanganisira kufumura zvirimwa zvemupunga kune tembiricha yepamusoro masikati/husiku (40°/30°C masikati/usiku). Koramu yega yega inomiririra iyo ± yakajairwa kukanganisa kwema data mashanu mapoinzi (n = 5). Makolamu anoteverwa nemabhii akasiyana anoratidza mutsauko wakakosha maererano nebvunzo yaTukey (P ≤ 0.05). Mavara ane chiratidzo chakaenzana anoratidza kuti chirevo hachina kukosha (≤ 0.05).
Principal component analysis (PCA) yakaratidza kuti zvipembenene zvakaongororwa pa55 DAE zvakatsanangura 66.1% yehutano hwehupenyu uye hwehupenyu hwehupenyu hwekupisa-kusimbisa rice miti inobatwa nekukura regulator spray (Fig. 3). Vectors vanomiririra zvinosiyana uye madotsi anomiririra chirimwa kukura regulators (GRs). Iwo mavector e gs, chlorophyll yemukati, yakanyanya kuwanda kwehuwandu hwePSII (Fv / Fm) uye biochemical parameters (TChl, MDA uye proline) ari padyo nemakona kune kwaakabva, zvichiratidza kuwirirana kwepamusoro pakati pehunhu hwepanyama hwezvirimwa nazvo. variable. Rimwe boka (V) raisanganisira madzinde emupunga anorimwa patembiricha yakanyatsonaka (AT) neF2000 zvirimwa zvinorapwa neCK neBA. Panguva imwecheteyo, ruzhinji rwemiti yakarapwa neGR yakagadzira boka rakasiyana (IV), uye kurapwa neGA muF2000 kwakagadzira boka rakasiyana (II). Kusiyana neizvi, mbesa dzemupunga dzakasimbiswa nekupisa (mapoka I uye III) pasina chero foliar spray ye phytohormones (ose genotypes aive SC) aive munzvimbo yakatarisana neboka V, zvichiratidza mhedzisiro yekupisa kwekushushikana pachirimwa physiology. .
Mufananidzo 3. Bigraphical analysis of the effects of joint heat stress (40°/30°C day/night) pamiti yemupunga miviri genotypes (F67 neF2000) pamazuva 55 mushure mekubuda (DAE). Kudimburira: AC F67, zvachose kutonga F67; SC F67, kupisa kudzvinyirira kuzvidzora F67; AUX F67, kupisa kunetseka + auxin F67; GA F67, kupisa kunetseka + gibberellin F67; CK F67, kupisa kunetseka + sero division BR F67, kupisa kunetseka + brassinosteroid. F67; AC F2000, zvachose kutonga F2000; SC F2000, Heat Stress Control F2000; AUX F2000, kupisa kunetseka + auxin F2000; GA F2000, kupisa kunetseka + gibberellin F2000; CK F2000, kushushikana kwekupisa + cytokinin, BR F2000, kushushikana kwekupisa + brass steroid; F2000.
Zvinhu zvakasiyana-siyana zvakadai se chlorophyll content, stomatal conductance, Fv/Fm ratio, CSI, MDA, RTI uye proline content zvinogona kubatsira kunzwisisa kugadziriswa kwemupunga genotypes uye kuongorora kukanganisa kweagronomic mazano pasi pekupisa kwekupisa (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Chinangwa chekuedza uku chaiva chekuongorora maitiro ekushandiswa kwezvigadziridzo zvina zvekukura pane physiological uye biochemical parameters yembesa yemupunga pasi pemamiriro ezvinhu akaoma ekupisa. Kuongororwa kwembesa inzira iri nyore uye inokurumidza yekuongorora panguva imwe chete yezvirimwa zvemupunga zvichienderana nehukuru kana mamiriro ezvivakwa zviripo (Sarsu et al. 2018). Zvigumisiro zvechidzidzo ichi zvakaratidza kuti kusanganiswa kwekupisa kunokonzera maitiro akasiyana-siyana ehupenyu uye biochemical mhinduro mune maviri mupunga genotypes, zvichiratidza maitiro ekugadzirisa. Mhedzisiro iyi inoratidzawo kuti foliar kukura regulator sprays (kunyanya cytokinins uye brassinosteroids) anobatsira mupunga kuchinjika kune yakaoma kupisa kusagadzikana sezvo favour inonyanya kubata gs, RWC, Fv/Fm ratio, photosynthetic pigments uye proline zvirimo.
Kushandiswa kwezvigadziriso zvekukura kunobatsira kuvandudza mamiriro emvura ezvirimwa zvemupunga pasi pekupisa kwekushushikana, izvo zvinogona kusanganiswa nekushushikana kwakanyanya uye kuderera kwekushisa kwechirimwa. Ichi chidzidzo chakaratidza kuti pakati pe "F2000" (inotapukira genotype) zvirimwa, zvirimwa zvemupunga zvakabatwa zvakanyanya neCK kana BR zvaive nepamusoro gs kukosha uye yakaderera PCT kukosha pane zvirimwa zvakabatwa neSC. Zvidzidzo zvakapfuura zvakaratidzawo kuti gs uye PCT ndeyechokwadi physiological zviratidzo zvinogona kuona mhinduro inogadziriswa yezvirimwa zvemupunga uye mhedzisiro yeagronomic mazano pakupisa kusagadzikana (Restrepo-Diaz naGarces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). Leaf CK kana BR inosimudzira g pasi pekunetseka nekuti aya mahomoni echirimwa anogona kukurudzira kuvhurika kwestomatal kuburikidza nekuita kwekuita nemamwe mamorekuru ekusaina akadai seABA (mutsigiri wekuvhara stomatal pasi pekunetseka kweabiotic) (Macková et al., 2013; Zhou et al., 2013). 2013). ) , 2014). Kuvhura kwestomatal kunosimudzira kutonhora kwemashizha uye kunobatsira kuderedza tembiricha yecanopy (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Nezvikonzero izvi, tembiricha yecanopy yemiti yemupunga yakapfapfaidzwa neCK kana BR inogona kunge yakadzikira pasi pekushushikana kwekupisa kwakasanganiswa.
Kunyanya kupisa kwekushisa kunogona kuderedza iyo photosynthetic pigment yemukati yemashizha (Chen et al., 2017; Ahammed et al., 2018). Muchidzidzo ichi, apo zvidyarwa zvemupunga zvaive pasi pekupisa kwekupisa uye zvisina kupfapfaidzwa nechero zvigadziriso zvekukura kwezvirimwa, photosynthetic pigments yaiwanzodzikira mune ese genotypes (Table 2). Feng et al. (2013) yakashumawo kuderera kukuru kweklorophyll yemukati mumashizha maviri egorosi genotypes akafumurwa nekupisa kunetseka. Kuratidzwa kune tembiricha yakakwira kazhinji kunokonzeresa kuderera kweklorophyll yemukati, izvo zvingave zvichikonzerwa nekudzikira kwechlorophyll biosynthesis, kuderedzwa kwepigment, kana mhedzisiro yavo yakasanganiswa pasi pekunetseka kwekupisa (Fahad et al., 2017). Nekudaro, zvirimwa zvemupunga zvakarapwa zvakanyanya neCK neBA zvakawedzera kuwanda kwemashizha photosynthetic pigments pasi pekupisa kwekushushikana. Migumisiro yakafanana yakataurwawo naJespersen and Huang (2015) uye Suchsagunpanit et al. (2015), uyo akaona kuwedzera kwemashizha chlorophyll yemukati zvichitevera kushandiswa kwezeatin uye epibrassinosteroid mahomoni mukupisa-kusimbisa bentgrass uye mupunga, zvichiteerana. Tsanangudzo inonzwisisika yekuti nei CK neBR vachikurudzira kuwedzera mashizha e chlorophyll pasi pekubatana kwekupisa kushushikana ndeyekuti CK inogona kuwedzera kutangwa kwekusimudzira kuderedzwa kwevanokurudzira kutaura (senge senescence-activating promoter (SAG12) kana HSP18 mutsigiri) uye kuderedza chlorophyll kurasikirwa mumashizha. , kunonoka mashizha senescence uye kuwedzera kumira kwechirimwa pakupisa (Liu et al., 2020). BR inogona kuchengetedza shizha chlorophyll uye kuwedzera mashizha chlorophyll yemukati ne activate kana kukonzeresa synthesis yema enzymes anobatanidzwa mu chlorophyll biosynthesis pasi pekunetseka mamiriro (Sharma et al., 2017; Siddiqui et al., 2018). Pakupedzisira, maviri phytohormones (CK uye BR) anokurudzirawo kutaura kwekupisa kunotyisa mapuroteni uye anovandudza akasiyana-siyana ekugadzirisa metabolic maitiro, akadai sekuwedzera chlorophyll biosynthesis (Sharma et al., 2017; Liu et al., 2020).
Chlorophyll a fluorescence parameters inopa nzira inokurumidza uye isiri-inoparadza iyo inogona kuongorora kushivirira kwezvirimwa kana kugadzirisa kune abiotic stress mamiriro (Chaerle et al. 2007; Kalaji et al. 2017). Maparamendi akadai seyeyero yeFv/Fm akashandiswa sezviratidzo zvekugadzirisa chirimwa kumamiriro ekushushikana (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Muchidzidzo ichi, zvidyarwa zveSC zvakaratidza hutsika hwakaderera hweiyi shanduko, kunyanya "F2000" zvirimwa zvemupunga. Yin et al. (2010) vakawanawo kuti Fv/Fm reshiyo yemashizha emupunga wepamusoro wakadzikira zvakanyanya patembiricha iri pamusoro pe35°C. Maererano naFeng et al. (2013), iyo yakaderera Fv/Fm reshiyo pasi pekupisa kushushikana kunoratidza kuti mwero wekunakidzwa kwesimba kubatwa uye kutendeuka nePSII reaction Center inoderedzwa, zvichiratidza kuti iyo PSII reaction centre inoparara mukushushikana kwekupisa. Kucherekedza uku kunotibvumira kugumisa kuti kuvhiringidzika mumidziyo yephotosynthetic kunonyanya kutaurwa mumhando dzakaoma (Fedearroz 2000) pane mumhando dzinopesana (Fedearroz 67).
Kushandiswa kweCK kana BR kazhinji kwakasimudzira kushanda kwePSII pasi pemamiriro akaoma ekupisa. Migumisiro yakafanana yakawanikwa na Suchsagunpanit et al. (2015), uyo akacherechedza kuti BR application yakawedzera kushanda kwePSII pasi pekupisa kunetseka mumupunga. Kumar et al. (2020) yakawanawo kuti zvirimwa zvechickpea zvakabatwa neCK (6-benzyladenine) uye zvakaiswa mukupisa kunetseka kwakawedzera chiyero cheFv/Fm, zvichigumisa kuti foliar application yeCK nekuita kuti zeaxanthin pigment cycle inosimudzira PSII chiitiko. Pamusoro pezvo, BR leaf spray yaifarira PSII photosynthesis pasi pemamiriro ekushushikana akabatana, zvichiratidza kuti kushandiswa kwephytohormone iyi kwakakonzera kuderera kwekuparara kwesimba rekukurudzira rePSII antennae uye kwakasimudzira kuunganidzwa kwediki kupisa kupisa mapuroteni mumakloroplast (Ogweno et al. 2008; Kothari naLachowitz). , 2021).
MDA uye zvinyorwa zveproline zvinowanzowedzera kana zvinomera zviri pasi pekunetseka kweabiotic zvichienzaniswa nemiti inokura pasi pemamiriro ezvinhu akanaka (Alvarado-Sanabria et al. 2017). Zvidzidzo zvakapfuura zvakaratidzawo kuti MDA uye mazinga eproline zviratidzo zvebiochemical zvinogona kushandiswa kunzwisisa maitiro ekugadzirisa kana kukanganisa kweagronomic maitiro mupunga pasi pemasikati kana usiku kupisa kwakanyanya (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al. . , 2021). Zvidzidzo izvi zvakaratidzawo kuti MDA uye proline zviri mukati zvaiwanzove zvakakwirira muzvirimwa zvemupunga zvakaratidzwa kune tembiricha yepamusoro husiku kana masikati, zvichiteerana. Zvisinei, kupfapfaidza foliar kweCK neBR kwakabatsira mukuderera kweMDA uye kuwedzera kwemazinga eproline, kunyanya mune inoshivirira genotype (Federroz 67). CK spray inogona kukurudzira kuwandisa kwe cytokinin oxidase / dehydrogenase, nekudaro ichiwedzera zviri mukati mekudzivirira kwemakemikari akadai se betaine uye proline (Liu et al., 2020). BR inosimudzira kuiswa kweosmoprotectants senge betaine, shuga, uye amino acids (kusanganisira yemahara proline), kuchengetedza cellular osmotic chiyero pasi pemamiriro mazhinji akashata ezvakatipoteredza (Kothari naLachowiec, 2021).
Crop stress index (CSI) uye hama yekushivirira indekisi (RTI) inoshandiswa kuona kana marapirwo ari kuongororwa anobatsira kurerutsa kushushikana kwakasiyana (abiotic uye biotic) uye kuve nemhedzisiro yakanaka pachirimwa physiology (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). CSI tsika dzinogona kubva ku0 kuenda ku1, inomiririra kusiri-kushushikana uye kusagadzikana mamiriro, zvichiteerana (Lee et al., 2010). Iwo CSI kukosha kwekupisa-kusimbiswa (SC) zvidyarwa kubva pa0.8 kusvika 0.9 (Mufananidzo 2B), zvichiratidza kuti zvirimwa zvemupunga zvakakanganiswa nekushushikana kwakasanganiswa. Nekudaro, kupfapfaidza foliar kweBC (0.6) kana CK (0.6) zvakanyanya kutungamirira mukudzikira kwechiratidzo ichi pasi pemamiriro ekushushikana kweabiotic zvichienzaniswa neSC mupunga zvirimwa. Mumiti yeF2000, RTI yakaratidza kuwedzera kwepamusoro pakushandisa CA (97.69%) uye BC (60.73%) zvichienzaniswa neSA (33.52%), zvichiratidza kuti izvi zvinomera kukura kwemiti zvinobatsirawo kuvandudza mhinduro yemupunga kune kushivirira kwekuumbwa. Overheat. Aya ma indices akarongedzerwa kugadzirisa mamiriro ekushushikana mumhando dzakasiyana. Chidzidzo chakaitwa naLee et al. (2010) yakaratidza kuti CSI yemhando mbiri dzekotoni pasi pekunetseka kwemvura yakadzikama yaive inosvika 0.85, nepo CSI tsika dzemhando dzakadiridzwa zvakanaka dzakabva ku0.4 kusvika 0.6, zvichigumisa kuti iyi index chiratidzo chekugadziriswa kwemvura yemhando. mamiriro ezvinhu anonetsa. Uyezve, Chavez-Arias et al. (2020) yakaongorora kushanda kwemaitiro ekugadzira elicitors senzira yakazara yekugadzirisa kushungurudzika muC. elegans miti uye yakaona kuti zvinomera zvakapfapfaidzwa nemishonga iyi yakaratidza yakakwirira RTI (65%). Kubva pane zviri pamusoro, CK neBR zvinogona kutorwa seagronomic mazano ane chinangwa chekuwedzera kushivirira kwemupunga kune yakaoma kupisa kunetseka, sezvo izvi zvinogadzirisa kukura kwezvirimwa zvinokurudzira mhinduro dzakanaka dzebiochemical uye physiological.
Mumakore mashoma apfuura, tsvakiridzo yemupunga muColombia yakatarisana nekuongorora genotypes inoshivirira kune yakakwira masikati kana husiku tembiricha uchishandisa physiological kana biochemical maitiro (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Nekudaro, mumakore mashoma apfuura, kuongororwa kwehunyanzvi hwekuita, hwehupfumi uye hune pundutso hwave huchiwedzera kukosha kupa zano rekubatanidza goho rekugadzirisa mhedzisiro yenguva dzakaoma dzekupisa kusagadzikana munyika (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021). Nokudaro, mhinduro dzehupenyu uye biochemical yemiti yemupunga kune yakaoma kupisa kunetseka (40 ° C zuva / 30 ° C usiku) yakaonekwa muchidzidzo ichi inoratidza kuti foliar spraying neCK kana BR inogona kuva nzira yakakodzera yekugadzirisa zvirimwa kuderedza migumisiro yakaipa. Mhedzisiro yenguva dzepakati kupisa kunetseka. Mishonga iyi yakavandudza kushivirira kweese mupunga genotypes (yakaderera CSI uye yakakwira RTI), zvichiratidza maitiro akajairwa muchirimwa physiological uye biochemical mhinduro pasi pekubatana kwekushushikana kwekupisa. Mhinduro huru yemiti yemupunga yaive kuderera kwemukati weGC, yakazara chlorophyll, chlorophylls α uye β uye carotenoids. Pamusoro pezvo, zvinomera zvinotambura nePSII kukuvadzwa (yakaderera chlorophyll fluorescence paramita senge Fv/Fm ratio) uye yakawedzera lipid peroxidation. Kune rumwe rutivi, apo mupunga wakabatwa neCK uye BR, izvi zvakashata zvakaderedzwa uye zvinyorwa zveproline zvakawedzera (Fig. 4).
Mufananidzo 4. Conceptual modhi yemhedzisiro yekusanganiswa kupisa kusagadzikana uye foliar chirimwa kukura regulator pfapfaidzo pamiti yemupunga. Miseve yakatsvuka uye yebhuruu inoratidza zvakaipa kana zvakanaka mhedzisiro yekudyidzana pakati pekupisa kunetseka uye foliar application yeBR (brassinosteroid) uye CK (cytokinin) pane zvepanyama uye biochemical mhinduro, zvichiteerana. gs: stomatal conductance; Total Chl: yakazara chlorophyll content; Chl α: chlorophyll β zviri mukati; Cx+c: carotenoid content;
Muchidimbu, mhinduro dzemuviri uye biochemical muchidzidzo ichi dzinoratidza kuti Fedearroz 2000 zvidyarwa zvemupunga zvinotapukirwa nenguva yekunetseka kwekupisa kupfuura Fedearroz 67 zvirimwa zvemupunga. Zvose zvinotungamirira kukura zvakaongororwa muchidzidzo ichi (auxins, gibberellins, cytokinins, kana brassinosteroids) zvakaratidza imwe dhigirii yekubatana kwekuderedza kushungurudzika. Nekudaro, cytokinin uye brassinosteroids yakakonzera zvirinani kuchinjika kwechirimwa sezvo ese ari maviri ekukura kwezvirimwa akawedzera chlorophyll content, alpha-chlorophyll fluorescence parameters, gs uye RWC zvichienzaniswa nezvirimwa zvemupunga pasina chero application, uye zvakare yakadzikira MDA zvirimo uye canopy tembiricha. Muchidimbu, tinogumisa kuti kushandiswa kwezvinomera zvinomera zvinogadzirisa (cytokinins uye brassinosteroids) chinhu chinobatsira pakugadzirisa mamiriro ekunetseka muzvirimwa zvemupunga zvinokonzerwa nekupisa kwakanyanya panguva yekupisa kwepamusoro.
Zvinyorwa zvepakutanga zvakaratidzwa muchidzidzo zvinosanganisirwa nechinyorwa, uye mimwe mibvunzo inogona kunangana nemunyori anoenderana.
Nguva yekutumira: Aug-08-2024